Metacommunity patterns of highly diverse stream midges: gradients,chequerboards, and nestedness,or is there only randomness?

Abstract.  1. Several non-random patterns in the distribution of species have been observed, including Clementsian gradients, Gleasonian gradients, nestedness, chequerboards, and evenly spaced gradients. Few studies have examined these patterns simultaneously, although they have often been studied in isolation and contrasted with random distribution of species across sites.
2. This study examined whether assemblages of chironomid midges exhibit any of the idealised distribution patterns as opposed to random distribution of species across sites within the metacommunity context in a boreal drainage system. Analyses were based on stream surveys conducted during three consecutive years. Analytical approaches included ordinations, cluster analysis, null models, and associated randomisation methods.
3. Midge assemblages did not conform to Clementsian gradients, which was evidenced by the absence of clearly definable assemblage types with numerous species exclusive to each assemblage type. Rather, there were signs of continuous Gleasonian variability of assemblage composition, as well as significant nested subset patterns of species distribution.
4. Midge assemblages showed only weak relationships with any of the measured environmental variables, and even these weak environmental relationships varied among years.
5. Midge assemblages did not appear to be structured by competition. This finding was somewhat problematic, however, because the two indices measuring co-occurrence provided rather different signs of distribution patterns. This was probably a consequence of how they actually measure co-occurrence.
6. Although midge assemblages did not show a perfect match with any of the idealised distribution patterns, they nevertheless showed a resemblance to the empirical patterns found previously for several plant and animal groups.     

Complex relationships between species niches and environmental heterogeneity affect species co-occurrence patterns in modelled and real communities

Species co-occurrence analysis is commonly used to assess how interspecific interactions dictate community assembly. Non-random co-occurrences, however, may also emerge from niche differences as well as environmental heterogeneity. The relationships between species co-occurrence patterns, environmental heterogeneity and species niches are not fully understood, due to complex interactions among them. To analyse the relationships among these patterns and processes, I developed synthetic community models and analysed a large dataset of tree species across the conterminous United States. Niche overlap and environmental heterogeneity had significant and contrasting effects on species co-occurrence patterns, in both modelled and real communities. Niche breadth, in turn, affected the effect sizes of both variables on species co-occurrence patterns. The effect of niche breadth on the relationship between co-occurrence and niche overlap was markedly consistent between modelled and real communities, while its effect on the relationship between co-occurrence and environmental heterogeneity was mostly consistent between real and modelled data. The results of this analysis highlight the complex and interactive effects of species niche overlap, niche breadth and environmental heterogeneity on species co-occurrence patterns. Therefore, inferring ecological processes from co-occurrence patterns without accounting for these fundamental characteristics of species and environments may lead to biased conclusions.     

Small mammal community structure in West Africa: a meta‐analysis using null models

Peer‐reviewed studies on the community structure of small mammals (Rodentia and Soricomorpha) from the Gulf of Guinea region of West Africa were reviewed. To detect nonrandom patterns in the various assemblages under study, the original datasets were re‐analysed using null models [two independent randomization algorithms (RA) for niche overlap] and Monte Carlo simulations. The total species richness in the countries considered in this review was 45 species for soricomorphs and 101 for rodents, and the studies reviewed here reported data for 53.3% of these soricomorph species and for 76.2% of these rodent species. Nonrandom habitat niche partitioning was rarely observed in both rodents and soricomorphs. Instead, aggregated use of habitat resources was frequently detected in both groups. Forest habitat was generally selected as aggregating resource type by small mammal assemblages. Thus, contrary to expectations, our review revealed little evidence for interspecific competition along the habitat niche axis in West African small mammals. However, it is possible that the aggregated use of the forest resource by small mammal species may be apparent, but that the various species partitioned space at a lower scale, for instance selecting different microhabitats. Interspecific competition appeared to be stronger in altered habitats, as predicted by previous studies on other forest organisms in West Africa.     

Fish abundance and habitat relationships in forest and grassland streams, northern Hokkaido, Japan

The relationships between habitat variables and population densities of masu salmon (Oncorhynchus masou), rosyface dace (Leuciscus ezoe), Siberian stone loach (Noemacheilus barbatulus) and wrinklehead sculpin (Cottus nozawae) were examined by data collected at 55 reaches in forest and grassland streams in northern Hokkaido, Japan. Regression analysis suggested that salmon and dace densities were affected by water temperature (negative for salmon, positive for dace) and structural habitat factors (woody debris for salmon, pools for dace). Salmon density was higher in forest reaches than in grassland reaches, whereas dace density was higher in grassland reaches, suggesting that the removal of riparian forest had raised water temperature and allowed upstream invasions by dace. In contrast to salmon and dace, neither the density of loach nor sculpin differed between the forest and grassland reaches. For their densities, a negative effect of each on the other was most important, suggesting a strong effect of interspecific competition between loach and sculpin on their distributions. However, regression models also suggested that substrate heterogeneity mediated the outcome of their interspecific competition. On the basis of the results, a scenario is predicted for a fish-assemblage change with a typical land-development process in Hokkaido, and the importance of leaving or restoring riparian buffer for conservation and restoration of stream habitat is emphasized.     

典型河床底质组成中底栖动物群落及多样性

底栖动物是河流生态系统中食物链的重要环节。通过对长江、黄河、东江和拒马河等河流野外调查和采样分析研究了河床底质组成对底栖动物群落结构的影响规律。研究结果发现,不同河床底质组成中的底栖动物结构差别很大,不同地理位置而相同底质条件和水力条件的河流底栖动物群落组成相似,说明河床底质是影响河流底栖动物群落结构的关键因素,受地理位置和大气候的影响不大;利用多项生物指标分析了不同河床底质组成中底栖动物群落的多样性,卵石河床且有水生植物生长的河流底栖动物物种组成最丰富,大河中沙质河床不稳定,未采集到底栖动物;不同底质类型河床中的优势种群亦不同。并分析了采样所得底栖动物物种数与采样面积之间的关系,符合前者随后者呈幂指数增加的规律,当实测采样面积为1～2m^2时物种数变化不大,建议一般情况下最小采样面积应为1m^2。     

长白山暗针叶林群落物种多样性维持机制

植物群落物种多样性维持机制一直是生态学研究的热点话题,其中生态位理论和中性理论是被普遍接受的两种理论观点,但是目前关于生态位理论和中性理论在群落物种多样性维持中的相对重要性还没有统一定论。基于长白山暗针叶林群落数据,采用单物种-面积关系模型探究特定树种对邻域物种丰富度的影响,并借助同质性和异质性泊松零模型检验其显著性。(1)群落水平上,在3—15 m空间尺度上,促进种占据优势地位,在>15 m空间尺度上,中性种逐渐取代促进种起主导作用,抑制种比例较低,并且随着空间尺度变化幅度不大。(2)物种水平上,采用同质性泊松零模型检验树种对邻域物种丰富度的影响,臭冷杉、花楷槭、青楷槭在0—20 m空间尺度上对邻域物种丰富度增加起促进作用,黄花落叶松、鱼鳞云杉在0—20 m空间尺度上抑制了邻域物种丰富度增加。花楸树、黑桦和硕桦在全部研究尺度上表现为中性种,髭脉槭、大青杨、红松等在不同研究尺度上表现为不同的作用效果。剔除了生境过滤作用的异质性泊松零模型检验结果与同质性泊松零模型结果差异不显著,表明研究样地内生境过滤作用对多样性格局形成影响不大,各树种间的相互作用对群落物种组成影响较大,进一步证明...     

Functional diversity and trait–environment relationships of stream fish assemblages in a large tropical catchment

1. The species composition of stream fish assemblages changes across the longitudinal fluvial gradient of large river basins. These changes may reflect both zonation in species distributions and environmental filtering of fish traits as stream environments change from the uplands to the lowlands of large catchments. Previous research has shown that taxonomic diversity generally increases in larger, lowland streams, and the River Continuum Concept, the River Habitat Template and other frameworks have provided expectations for what functional groups of fishes should predominate in certain stream types. However, studies addressing the functional trait composition of fish assemblages across large regions are lacking, particularly in tropical river basins. 2. We examined functional trait–environment relationships and functional diversity of stream fish assemblages in the Río Grijalva Basin in southern Mexico. Traits linked to feeding, locomotion and life history strategy were measured in fishes from streams throughout the catchment, from highland headwaters to broad, lowland streams. Relationships between functional traits and environmental variables at local and landscape scales were examined using multivariate ordination, and the convex hull volume of trait space occupied by fish assemblages was calculated as a measure of functional diversity. 3. Although there were a few exceptions, functional diversity of assemblages increased with species richness along the gradient from uplands to lowlands within the Grijalva Basin. Traits related to swimming, habitat preference and food resource use were associated with both local (e.g. substratum type, pool availability) and landscape‐scale (e.g. forest cover) environmental variables. 4. Along with taxonomic structure and diversity, the functional composition of fish assemblages changed across the longitudinal fluvial gradient of the basin. Trait–environment relationships documented in this study partially confirmed theoretical expectations and revealed patterns that may help in developing a better understanding of general functional responses of fish assemblages to environmental change.     

Seasonal modifications of longitudinal distribution patterns within a stream: Interspecific interactions in the niche overlap zones of two Ephemera mayflies

Niche differentiation between closely related species leads to differentiation of their habitats. Segregation based on slight differences in environmental factors, that is niche differentiation on the microhabitat scale, allows more species to inhabit a certain geographic space. Therefore, such fine scale niche differentiation is an important factor in the support of species diversity. In addition, niche differentiation on the microhabitat scale and/or the differentiation of breeding seasons can be considered typical mechanisms that facilitate multispecies’ co‐existence. In this study, sister species (Commonly, Ephemera japonica inhabit at upstream region and Ephemera strigata inhabit at middle stream region), which often coexist in the upper to middle reaches of river systems of the Japanese Islands, were targeted and the following aspects were investigated. First, differences in habitat preference and interspecific differences in flow distribution patterns on a geographically fine scale were tracked in detail. Subsequently, the temporal transitions of their distribution patterns were investigated in detail and seasonal changes were investigated. Finally, we thoroughly investigated the disappearance of nymphs of each species from the river due to emergence affected the distribution of each species (by conducting daily emergence surveys). Combining results of these multiple studies also suggested that there may be spatiotemporal interspecial interaction between these two species within/around their overlapping regions. Traditionally, the longitudinal distribution pattern of these two Ephemera mayflies has been thought to be established based on a difference in habitat preferences, but this study revealed that the interspecific interaction between the two species also plays an important role. This study provides new insights into species diversity and distribution pattern formation in river‐dwelling species.     

Habitat segregation in four species of adult tiger beetles (Coleoptera: Cicindelidae)

ABSTRACT.

• 1 Habitat segregation in four species of tiger beetles, Cicindela cancellata Dejean, C.cardoni Fleutians, C.minuta Olivier and C.sumatrensis Herbst in a river bank ecosystem was studied in dry and wet seasons.
• 2 The four species segregated distinctly along the river beds into separate habitats, with occasional overlapping in both the seasons.
• 3 Among the habitat characteristics considered, vegetation, soil moisture and available prey-size were found to be important in species segregation.
• 4 D ²-analysis showed that the habitat preferences of C.cardoni and C.cancellata were closer to that of C.sumatrensis. The habitat of C.minuta was distinctly separated.
• 5 There was a positive correlation between the mandible length of each tiger beetle species and the length of prey captured.
• 6 Variance in mandible length within species was related to prey-length distribution pattern and to variance in habitats.

     

Local‐scale spatial dynamics of ants in a temperate agroecosystem

At the local scale, spatial aggregations in ant distribution are often thought to be driven by competitive interactions among dominant ant species, although niche preferences and habitat heterogeneity might also lead to patchiness. Nevertheless, competitive interactions might be particularly important in agroecosystems that are structurally more homogeneous than natural habitats. The spatial patterns of ants in two Australian vineyards were investigated by intensive pitfall trapping to examine if non‐random patterns occur and whether these might be the result of competitive species interactions as well as the influence of woody vegetation adjacent to the vineyards. Null model analyses suggested competitive species interactions within ant assemblages that might have been driven by numerically dominant species, even though both positive and negative associations between these were found. Consistent spatial aggregations indicated significant spatial overlap in distributions of some species. Such overlap suggests that potential coexistence might be attributed to temporal partitioning or differences in foraging strategies. The presence of woody vegetation had a marked influence on ant assemblage structure and competitive interactions, and might facilitate coexistence by increasing resource heterogeneity. The implications of these findings for sampling strategies and ecological processes within vineyards are discussed.     

Spatial Patterns of Aggression, Territory Size, and Harem Size in Five Sympatric Caribbean Parrotfish Species

Territorial behaviour is a conspicuous determinant of social organisation in many reef fishes including parrotfishes. Most parrotfish studies in the Caribbean have focused on the species Scarus iserti and Sparisoma viride over limited ranges of reef habitat. By contrast, our study has included all common parrotfishes in Belize (Sc. iserti, Sp. viride, Sparisoma aurofrenatum, Sparisoma chrysopterum, and Sparisoma rubripinne) at three sites with different physical and biotic conditions and a wide range of fish densities. Density in Sc. iserti was strongly positively correlated to substrate rugosity. In contrast, densities of Sp. chrysopterum and Sp. rubripinne were unrelated to rugosity and territories were large. Territory size was smallest in Sc. iserti (mean areas at the three sites ranged from 41 to 120m²) and largest in Sp. rubripinne (ranged from 168 to 1400m²). All species except Sp. chrysopterum exhibited significantly larger territories where density was low as suggested by territory theory. Territory size decreased rapidly with increasing density of competitors. Patterns of harem size differed between two groups of parrotfishes. (1) Sc. iserti, Sp. viride, and Sp. aurofrenatum exhibited an expected positive correlation with territory size. (2) Harem size was smaller in Sp. rubripinne and Sp. chrysopterum, and showed no spatial pattern. Aggression in Sp. viride and Sc. iserti was directed entirely towards intraspecifics and positively density dependent. Interspecific interactions accounted for only 10% of observations and were recorded exclusively whilst following Sp. chrysopterum, Sp. rubripinne, and, to a lesser extent, Sp. aurofrenatum. A meta analysis of species interactions suggested that intraspecific interactions were most common where overall fish density was greatest and conversely, interspecific interactions occurred more often at lower densities. This may suggest that the economic defensibility of territories is largely confined to intraspecifics where density is greatest. Most (62%) of the interspecific interactions comprised Sp. rubripinne chasing the smaller species Sp. chrysopterum, suggesting that territorial behaviour has at least some non-reproductive origin and may therefore be associated with either food or shelter. It is feasible that at such low population densities, it is economically feasible for Sp. rubripinne to defend against intraspecifics and Sp. chrysopterum. Social behaviour in Sp. chrysopterum and Sp. rubripinne, and to a lesser extent Sp. aurofrenatum, differs to that of Sc. iserti and Sp. viride which conform to existing theories of social behaviour in reef fish.     

No short‐term change in avian assemblage following removal of Yellow‐throated Miner (Manorina flavigula) colonies

The overabundance of Yellow‐throated Miner (Manorina flavigula) has been shown to negatively affect the abundance and richness of small birds in areas they occupy, leading to homogenization of the avifauna across the fragmented landscape. In this study, we took advantage of a planned management cull to ask the question, does the removal of Yellow‐throated Miner colonies cause an immediate change in avian species richness and abundance? This cull was undertaken around the Bronzewing Flora and Fauna Reserve (north‐western Victoria, Australia) in order to protect a resident population of endangered Black‐eared Miner (M. melanotis) from hybridization. We conducted avian surveys along roadsides surrounding the reserve at Yellow‐throated Miner colonies (n = 6), control sites with no miners (n = 7), and where colonies were removed (n = 3). We found that the cull was followed by only a very modest increase in the species richness and abundance of small birds, with no significant effects on avian assemblage overall. This result contrasts with far more dramatic increases following culls of other species of miner. Sites where miners were removed were not depauperate of other species prior to the cull, which could have been due to a combination of proximity to refuge for small birds in a neighbouring reserve or the low numbers of miners that made up each culled colony. This study highlights that assumed effects of a management action may be highly dependent upon spatial and temporal context.     

Relationships between stream macroinvertebrates and environmental variables at multiple spatial scales

1. Aquatic communities are structured by multiple forces, and identifying the driving factors over multispatial scales is an important research issue. The East Asian monsoon region is globally one of the richest environments in terms of biodiversity, and is undergoing rapid human development, yet the river ecosystems in this region have not been well studied. We applied a hierarchical framework to incorporate regional and local environmental effects on stream macroinvertebrate communities in this region. The knowledge gained is expected to improve the understanding of the importance of spatial scale on regional and local diversity in the East Asian monsoon region. 2. A national data set of benthic macroinvertebrates and environmental variables (geographical, land‐use, hydrological, substratum and physicochemical elements) in Korean rivers was used to determine the habitat preferences of macroinvertebrates. 3. Latitude, proportion of forest coverage, riffle habitat, silt substratum and temperature were the most important determinants for the ordinations of macroinvertebrate communities in each category evaluated by canonical correspondence analysis (CCA). The optimal habitats for stream macroinvertebrates are not the same for all species, and overall community metrics and abundance of sensitive species tended to be lower in open agricultural and urban streams than in forested streams. The sensitivity of mayflies and stoneflies to anthropogenic disturbances implicated them as good indicators to assess the effects of urban and agricultural activities. 4. A partial CCA was used to evaluate the relative importance of macrohabitat and microhabitat variables on community composition at three spatial scales (whole country, the large Han River basin and two small sub‐basins in the lowlands and highlands). The majority of community variation (17–22% for each environmental element) was explained by macrohabitat variables at the regional spatial scale. In contrast, large proportions (15–18%) were explained by microhabitat variables at the local spatial scale. 5. Our findings indicate that the relative importance of habitat scales should be determined by geographical size and that comprehensive understanding of multispatial scale patterns can be important for implementing sound biodiversity conservation programmes.     

Climate‐change winners and losers: stream macroinvertebrates of a submontane region in Central Europe

1. Freshwater ecosystems will be profoundly affected by global climate change, especially those in mountainous areas, which are known to be particularly vulnerable to warming temperatures. We modelled impacts of climate change on the distribution ranges of 38 species of benthic stream macroinvertebrates from nine macroinvertebrate orders covering all river zones from the headwaters to large river reaches. 2. Species altitudinal shifts as well as range changes up to the year 2080 were simulated using the A2a and B2a Intergovernmental Panel on Climate Change climate‐warming scenarios. Presence‐only species distribution models were constructed for a stream network in Germany’s lower mountain ranges by means of consensus projections of four algorithms, as implemented in the BIOMOD package in R (GLM, GAM, GBM and ANN). 3. Species were predicted to shift an average of 122 and 83 m up in altitude along the river continuum by the year 2080 under the A2a and B2a climate‐warming scenarios, respectively. No correlation between altitudinal shifts and mean annual air temperature of species’ occurrence could be detected. 4. Depending on the climate‐warming scenario, most or all (97% for A2a and 100% for B2a) of the macroinvertebrate species investigated were predicted to survive under climate change in the study area. Ranges were predicted to contract for species that currently occur in streams with low annual mean air temperatures but expand for species that inhabit rivers where air temperatures are higher. 5. Our models predict that novel climate conditions will reorganise species composition and community structure along the river continuum. Possible effects are discussed, including significant reductions in population size of headwater species, eventually leading to a loss of genetic diversity. A shift in river species composition is likely to enhance the establishment of non‐native macroinvertebrates in the lower reaches of the river continuum.     

Lack of habitat segregation and no interspecific territoriality in three syntopic cryptic species of the golden‐spectacled warblers Phylloscopus (Seicercus) burkii complex

It is widely accepted that exploitative competition prevents the coexistence of any two or more closely‐related species unless differences exists in their ecological niches and resource use. In sibling bird species, exploitative competition is reduced mainly by spatial segregation of competing species. Spatial segregation can be achieved in two basic ways: by using different habitats or microhabitats that each species is predominately exploiting, or by interspecific territoriality. To our knowledge, either habitat segregation or interspecific territoriality or both have been found in all dyads and groups of sympartic sibling bird species studied so far. In this study, we investigated the coexistence of three cryptic sibling species of the golden‐spectacled warblers Phylloscopus burkii complex in Hunan Province, China: Ph. tephrocephalus, Ph. omeiensis and Ph. valentini. We analyzed their habitat preferences, and spatial interrelations between the species. In order to analyze either presence or absence of interspecific territoriality, we additionally performed con‐ and heterospecific playback experiments. Contrary to expectations, we found no evidences for either habitat segregation or interspecific territoriality in these three species.